Postnatal development of fiber type composition in rabbit jaw and leg muscles.

We examined the difference in fiber type composition and cross-sectional areas during postnatal development in male rabbit jaw muscles and compared these with changes in leg muscles. The myosin heavy chain (MyHC) content of the fibers was determined by immunohistochemistry. No fiber type difference was found between the jaw muscles in 20-week-old rabbits. However, the way this adult fiber type composition was reached differed between the muscles. The deep temporalis, medial pterygoid, and superficial masseter displayed an increase in alpha fibers during early and a decrease during late postnatal development. Other jaw muscles displayed an increase in alpha fibers during early development only. In contrast, alpha fibers were not found in the soleus, in which fiber type changes were completed at week 4. The gastrocnemius muscle did not change its fiber type composition. Initially, fibers in jaw-opening muscles had larger cross-sectional areas than in other muscles, but they increased less during development. Although there were no large differences in the fiber type composition of muscles in young adult rabbits, large differences were found in the jaw muscles, but not in the leg muscles, during development. In part, these developmental changes in fiber percentages within the jaw muscles can be explained by functional modifications in this muscle group. In the present study, the deep temporalis, medial pterygoid, and superficial masseter showed the most dramatic percent changes in fibers during postnatal development.

Is fiber-type composition related to daily jaw muscle activity during postnatal development?

AIM: Muscles containing large numbers of slow-contracting fibers are generally more active than muscles largely composed of fast fibers. This relationship between muscle activity and phenotype suggests that (1) changes in fiber-type composition during postnatal development are accompanied by changes in daily activity and (2) individual variations in fiber-type composition are related to similar variations in daily muscle activity. METHODS: The masseter and digastric muscles of 23 New Zealand White rabbits (young, juvenile and adult) were examined for their phenotype (myosin heavy chain content) and their daily activity (total daily number of activity bursts). RESULTS: During development, the masseter showed a strong increase in the number of fast-type fibers compared to the number of slow-type fibers. During development, also the number of powerful bursts in the masseter increased. The digastric showed no significant changes in fiber types or burst numbers. Within each muscle, across individual animals, no significant correlations (R < 0.70) were found between any of the fiber types and daily burst numbers in any of the age groups. CONCLUSIONS: The results suggest that activity-related influences are of relatively minor importance during development and that other factors are dominant in determining fiber-type composition.

Biomechanical effect of mineral heterogeneity in trabecular bone.

Due to daily loading, trabecular bone is subjected to deformations (i.e., strain), which lead to stress in the bone tissue. When stress and/or strain deviate from the normal range, the remodeling process leads to adaptation of the bone architecture and its degree of mineralization to effectively withstand the sustained altered loading. As the apparent mechanical properties of bone are assumed to depend on the degree and distribution of mineralization, the goal of the present study was examine the influences of mineral heterogeneity on the biomechanical properties of trabecular bone in the human mandibular condyle. For this purpose nine right condyles from human dentate mandibles were scanned and evaluated with a microCT system. Cubic regional volumes of interest were defined, and each was transformed into two different types of finite element (FE) models, one homogeneous and one heterogeneous. In the heterogeneous models the element tissue moduli were scaled to the local degree of mineralization, which was determined using microCT. Compression and shear tests were simulated to determine the apparent elastic moduli in both model types. The incorporation of mineralization variation decreased the apparent Young's and shear moduli by maximally 21% in comparison to the homogeneous models. The heterogeneous model apparent moduli correlated significantly with bone volume fraction and degree of mineralization. It was concluded that disregarding mineral heterogeneity may lead to considerable overestimation of apparent elastic moduli in FE models.

Spatial orientation in bone samples and Young's modulus.

Bone mass is the most important determinant of the mechanical strength of bones, and spatial structure is the second. In general, the spatial structure and mechanical properties of bones such as the breaking strength are direction dependent. The mean intercept length (MIL) and line frequency deviation (LFD) are two methods for quantifying directional aspects of the spatial structure of bone. Young's modulus is commonly used to describe the stiffness of bone, which is also a direction-dependent mechanical property. The aim of this article is to investigate the relation between MIL and LFD on one hand and Young's modulus on the other. From 11 human mandibular condyles, 44 samples were taken and scanned with high-resolution computer tomography equipment (micro-CT). For each sample the MIL and LFD were determined in 72602 directions distributed evenly in 3D space. In the same directions Young's modulus was determined by means of the stiffness tensor that had been determined for each sample by finite element analysis. To investigate the relation between the MIL and LFD on one hand and Young's modulus on the other, multiple regression was used. On average the MIL accounted for 69% of the variance in Young's modulus in the 44 samples and the LFD accounted for 72%. The average percentage of variance accounted for increased to 80% when the MIL was combined with the LFD to predict Young's modulus. Obviously MIL and LFD to some extent are complementary with respect to predicting Young's modulus. It is known that directional plots of the MIL tend to be ellipses or ellipsoids. It is speculated that ellipsoids are not always sufficient to describe Young's modulus of a bone sample and that the LFD partly compensates for this.

Regional variations in mineralization and strain distributions in the cortex of the human mandibular condyle.

The strain (i.e. deformation) history influences the degree of mineralization of cortical bone (DMB) as well as its osteonal microstructure. This study aimed to examine the relationships of stress and strain distributions with the variations in DMB and the osteonal orientations in the cortical bone of the human mandibular condyle. It was hypothesized that strains are inversely proportional to local DMB and that the principal strains are oriented parallel to the osteons. To test this, ten human mandibular condyles were scanned in a microCT system. Finite element models were created in order to simulate static clenching. Within each condyle, 18 volumes of interest were selected to analyze regional differences in DMB, stress and strains. Subchondral bone showed a lower equivalent strain (2652+/-612 muepsilon) as compared to the anterior (p=0.030) and posterior cortex (p=0.007) and was less mineralized. Contrary to our hypothesis, the results show that strains correlated positively with regional variations in DMB (r=0.750, p<0.001). In the anterior and the posterior cortex, the first principal strain was parallel to the cortical surface and oriented supero-inferiorly with a fan-like shape. In subchondral bone, the first and the second principal strain were parallel to the surface and oriented antero-posteriorly and medio-laterally, respectively. It was concluded that the strain distributions, by themselves, cannot explain the regional differences found in DMB. In agreement with our second hypothesis, the orientation of the osteonal network of the mandibular condyle was closely related to the strain orientations. The results of this study suggest that the subchondral and the cortical bone are structured to ensure an optimal load distribution within the mandibular condyle and have a different mechanical behaviour. Subchondral bone plays a major role in the transmission of the strains to the anterior and posterior cortex, while these ensure an optimal transmission of the strains within the condylar neck and, eventually, to the mandibular ramus.

Consequences of viscoelastic behavior in the human temporomandibular joint disc.

The consequences of the viscoelastic behavior of the temporomandibular joint disc were analyzed in simulated jaw open-close cycles. It was hypothesized that viscoelasticity helps protect the underlying bone, while augmenting the smoothness of articular movements. Simulations were performed with a dynamic model of the masticatory system, incorporating the joints' cartilaginous structures as Finite Element Models. A non-linear viscoelastic material model was applied for the disc. The apparent stiffness of the disc to principal stress was largest when the jaw was closed, whereas, with the Von Mises' stress, it appeared largest when the jaw was open. The apparent stiffnesses appeared to be dependent on both the speed of the movements and the presence of a resistance between the teeth. It was concluded that the disc becomes stiffer when load concentrations can be expected. During continued cyclic motion, it softens, which favors smoothness of joint movement at the cost of damage prevention.

Variations in mineralization affect the stress and strain distributions in cortical and trabecular bone.

The mechanical properties of bone depend largely on its degree and distribution of mineralization. The present study analyzes the effect of an inhomogeneous distribution of mineralization on the stress and strain distributions in the human mandibular condyle during static clenching. A condyle was scanned with a micro-CT scanner to create a finite element model. For every voxel the degree of mineralization (DMB) was determined from the micro-CT scan. The Young's moduli of the elements were calculated from the DMB using constant, linear, and cubic relations, respectively. Stresses, strains, and displacements in cortical and trabecular bone, as well as the condylar deformation (extension along the antero-posterion axis) and compliance were compared. Over 90% of the bone mineral was located in the cortical bone. The DMB showed large variations in both cortical bone (mean: 884, SD: 111 mg/cm(3)) and trabecular bone (mean: 738, SD: 101 mg/cm(3)). Variations of the stresses and the strains were small in cortical bone, but large in trabecular bone. In the cortical bone an inhomogeneous mineral distribution increased the stresses and the strains. In the trabecular bone, however, it decreased the stresses and increased the strains. Furthermore, the condylar compliance remained relatively constant, but the condylar deformation doubled. It was concluded that neglect of the inhomogeneity of the mineral distribution results in a large underestimation of the stresses and strains of possibly more than 50%. The stiffness of trabecular bone strongly influences the condylar deformation. Vice versa, the condylar deformation largely determines the magnitude of the strains in the trabecular bone.

Viscoelastic material model for the temporomandibular joint disc derived from dynamic shear tests or strain-relaxation tests.

Viscoelastic material models for the temporomandibular joint disc, based upon strain relaxation, were considered to underestimate energy absorption for loads with time constants beyond the relaxation time. Therefore, the applicability of a material model that takes the viscous behavior at a wide range of frequencies into account was assessed. To that purpose a non-linear multi-mode Maxwell model was tested in cyclic large-strain compression tests. Its material constants were approximated from dynamic small-strain shear deformation tests. The storage and loss moduli as obtained from a disc sample could be approximated with a four-mode Maxwell model. In simulated large-strain compression tests it behaved similarly as observed from the experimental tests. The underestimation of energy dissipation, as obtained from a single-mode Maxwell model was considerably reduced, especially for deformations with a higher strain rate. Furthermore, in contrast to the latter it was able to predict the increase of the stress amplitude with the compression frequency much better. In conclusion, the applied four-mode Maxwell model, based upon dynamic shear tests, was considered more suitable to predict higher frequency viscoelastic response, for instance during shock absorption, than a model based upon strain-relaxation.

Daily activity of the rabbit jaw muscles during early postnatal development.

Early postnatal development of the jaw muscles is characterized by the transition from suckling to chewing behavior. As chewing develops the jaw closing muscles become more powerful compared with the jaw openers. These changes are likely to affect the amount of daily muscle activity. Therefore, the purpose of this study was to characterize for a jaw opener (digastric) and jaw closer (masseter) the total duration of daily muscle activity (i.e. the duty time), and the daily burst numbers and lengths during early postnatal development. Using radiotelemetry the activity of these muscles was recorded in 10 young New Zealand White rabbits between three and eight weeks of age. Fiber-type composition was analyzed at eight weeks of age by determining the myosin heavy chain content of the fibers. During postnatal development both muscles showed no significant decrease or increase in their daily activity. However, the interindividual variation of the duty time and burst number significantly decreased. There were no significant differences between the digastric and masseter except for the most powerful activities at eight weeks of age, where the masseter showed a significantly higher duty time and burst number than the digastric. The masseter contained a higher number of slow-type fibers expressing myosin heavy chain-I and myosin heavy chain-cardiac alpha than the digastric. The present results suggest that the amount of jaw muscle activation is already established early during postnatal development, before the transition from suckling to chewing behavior. This amount of activation seems to be related to the number of slow-type fibers.

Duty time of rabbit jaw muscles varies with the number of activity bursts.

The relative duration of muscle activity during a specified period (duty time) varies depending on activity level and time of the day. Since both the number and the length of activity bursts contribute to the duty time, it was hypothesized that these variables would show intra-day variations similar to those of the duty time. To test this, we determined duty times, burst numbers, and burst lengths per hour, in relation to multiple activity levels, in a 24-hour period of concurrent radio-telemetric long-term electromyograms of various rabbit jaw muscles. The marked intra-day variation of the burst number resembled that of the duty time in all muscles, and was in contrast to the relatively invariable mean burst length. Furthermore, the duty times were more highly correlated with the number than with the length of bursts at all activity levels. Thus, the variation of the duty time in rabbit jaw muscles is caused mainly by changes in burst numbers.

Structural and mechanical properties of mandibular condylar bone.

The trabecular bone of the mandibular condyle is structurally anisotropic and heterogeneous. We hypothesized that its apparent elastic moduli are also anisotropic and heterogeneous, and depend on trabecular density and orientation. Eleven condyles were scanned with a micro-CT system. Volumes of interest were selected for the construction of finite element models. We simulated compressive and shear tests to determine the principal mechanical directions and the apparent elastic moduli. Compressive moduli were relatively large in directions acting in the sagittal plane, and small in the mediolateral direction. The degree of mechanical anisotropy ranged from 4.7 to 10.8. Shear moduli were largest in the sagittal plane and smallest in the transverse plane. The magnitudes of the moduli varied with the condylar region and were proportional to the bone volume fraction. Furthermore, principal mechanical direction correlated significantly with principal structural direction. It was concluded that variation in trabecular structure coincides with variation in apparent mechanical properties.

Dynamic compressive properties of the mandibular condylar cartilage.

The mandibular condylar cartilage plays an important role as a stress absorber during function. However, relatively little information is available on its dynamic properties under compression. We hypothesized that these properties are region-specific and depend on loading frequency. To characterize the viscoelastic properties of the condylar cartilage, we performed dynamic indentation tests over a wide range of loading frequencies. Ten porcine mandibular condyles were used; the articular surface was divided into 4 regions, anteromedial, anterolateral, posteromedial, and posterolateral. The dynamic complex, storage, and loss moduli increased with frequency, and these values were the highest in the anteromedial region. Loss tangent decreased with frequency from 0.68 to 0.17, but a regional difference was not found. The present results suggest that the dynamic compressive modulus is region-specific and is dependent on the loading frequency, which might have important implications for the transmission of load in the temporomandibular joint.

Passive resistance increases differentially in various jaw displacement directions.

OBJECTIVES: In the present study, the passive resistance of the human jaw system was quantified in relation to the three-dimensional jaw displacement and the Posselt-envelope, using both in vivo measurements and computer simulation. METHODS: In eight subjects, the jaw was passively displaced with a step-wise increasing force in three orthogonal directions. Muscle relaxation was monitored using electromyography (EMG) with visual feedback. A biomechanical model of an average human system was used to examine the contributions of the jaw muscles. RESULTS: The largest excursion was found for the vertical direction. Protrusive and lateral directions were more restricted. In protrusive and lateral directions, the jaw could generally move beyond the Posselt-envelope. The stiffness of the jaw increased with proceeding jaw displacement in all directions. The stiffness was larger in the protrusive direction than in the vertical and lateral directions. The model's predictions of stiffness were comparable to the in vivo measurements. However, in protrusive direction, the maximum jaw displacement was larger than in vivo. The estimated passive muscle forces showed that vertical displacement was mainly restricted by the complete group of closing muscles, while protrusive and lateral jaw displacement was restricted by selective individual muscles. CONCLUSIONS: The human jaw system has larger motion range in the protrusive and lateral directions than can be exploited by active muscle use. Stiffness of jaw displacement is higher in the protrusive direction compared to the vertical and lateral directions.

[Transient paresis after mandibular block anaesthesia]. / Parese na mandibulaire blokanesthesie.

Mandibular block anaesthesia is frequently used in dentistry. The technique is more susceptible to failure than local anaesthesia by infiltration. Two cases of patients who were afflicted by transient paresis of (branches of) the facial and vagus nerves after mandibular block anaesthesia have been analysed. With reference to both these cases advice is given on how to react in case of a misdirected mandibular block anaesthesia.

The effect of bone loss on rod-like and plate-like trabeculae in the cancellous bone of the mandibular condyle.

Bone loss may affect the structure of cancellous bone. But its effect on trabeculae with different characteristics, like rods and plates, is not accurately known. This study analyzes the effect of bone loss on individual rod-like and plate-like trabeculae. 94 specimens were obtained from mandibular condyles from both dentate and edentate humans and scanned with a micro-CT scanner. The bone volume fraction (BV/TV) of these specimens ranged from 7% to 30%. Next, the rod-like and plate-like trabeculae were identified with an especially developed algorithm. Plate volume fraction (PV/TV), rod volume fraction (RV/TV), plate thickness, rod thickness, number of plates, and number of rods were determined. In individual specimens, the thickness of the rods ranged from 40 mum to 180 mum, while the thickness of the plates ranged from 40 mum to 300 mum. In every specimen, the thickness of the plates was larger than the thickness of the rods. Statistical analysis revealed that PV/TV was proportional (r = 0.98, P < 0.001) and RV/TV inversely proportional (r = -0.45, P < 0.001) to BV/TV. Also the thickness of the plates correlated with BV/TV (r = 0.62, P < 0.001), while the thickness of the rods (mean = 90 mum, SD = 7 mum) remained constant (r = -0.09, P = 0.378). A four-fold reduction of the bone volume fraction was accompanied by a three-fold reduction of the number of plates and a 40% reduction of their thickness, but also by a three-fold increase in the number of rods and RV/TV. It was concluded that the effect of bone loss on plate-like trabeculae was opposite to its effect on rod-like trabeculae. Remarkably, the thickness of the rods (90 mum) was independent of the bone volume fraction. This suggests that there is a minimal thickness for trabeculae.

Fiber-type composition of the human jaw muscles--(part 1) origin and functional significance of fiber-type diversity.

This is the first of two articles on the fiber-type composition of the human jaw muscles. The present article discusses the origin of fiber-type composition and its consequences. This discussion is presented in the context of the requirements for functional performance and adaptation that are imposed upon the jaw muscles. The human masticatory system must perform a much larger variety of motor tasks than the average limb or trunk motor system. An important advantage of fiber-type diversity, as observed in the jaw muscles, is that it optimizes the required function while minimizing energy use. The capacity for adaptation is reflected by the large variability in fiber-type composition among muscle groups, individual muscles, and muscle regions. Adaptive changes are related, for example, to the amount of daily activation and/or stretch of fibers. Generally, the number of slow, fatigue-resistant fibers is relatively large in muscles and muscle regions that are subjected to considerable activity and/or stretch.

Fiber-type composition of the human jaw muscles--(part 2) role of hybrid fibers and factors responsible for inter-individual variation.

This is the second of two articles about fiber-type composition of the human jaw muscles. It reviews the functional relationship of hybrid fibers and the adaptive properties of jaw-muscle fibers. In addition, to explain inter-individual variation in fiber-type composition, we discuss these adaptive properties in relation to environmental stimuli or perturbations. The fiber-type composition of the human jaw muscles is very different from that of limb and trunk muscles. Apart from the presence of the usual type I, IIA, and IIX myosin heavy-chains (MyHC), human jaw-muscle fibers contain MyHCs that are typical for developing or cardiac muscle. In addition, much more frequently than in limb and trunk muscles, jaw-muscle fibers are hybrid, i.e., they contain more than one type of MyHC isoform. Since these fibers have contractile properties that differ from those of pure fibers, this relatively large quantity of hybrid fibers provides a mechanism that produces a very fine gradation of force and movement. The presence of hybrid fibers might also reflect the adaptive capacity of jaw-muscle fibers. The capacity for adaptation also explains the observed large inter-individual variability in fiber-type composition. Besides local influences, like the amount of muscle activation and/or stretch, more general influences, like aging and gender, also play a role in the composition of fiber types.

Combined finite-element and rigid-body analysis of human jaw joint dynamics.

The jaw joint plays a crucial role in human mastication. It acts as a guidance for jaw movements and as a fulcrum for force generation. The joint is subjected to loading which causes tensions and deformations in its cartilaginous structures. These are assumed to be a major determinant for development, maintenance and also degeneration of the joint. To analyze the distribution of tensions and deformations in the cartilaginous structures of the jaw joint during jaw movement, a dynamical model of the human masticatory system has been constructed. Its movements are controlled by muscle activation. The articular cartilage layers and articular disc were included as finite-element (FE) models. As this combination of rigid-body and FE modeling had not been applied to musculoskeletal systems yet, its benefits and limitations were assessed by simulating both unloaded and loaded jaw movements. It was demonstrated that joint loads increase with muscle activation, irrespective of the external loads. With increasing joint load, the size of the stressed area of the articular surfaces was enlarged, whereas the peak stresses were much less affected. The results suggest that the articular disc enables distribution of local contact stresses over a much wider area of the very incongruent articular surfaces by transforming compressive principal stress into shear stress.

Myosin heavy chain composition in human masticatory muscles by immunohistochemistry and gel electrophoresis.

In this study we compared the immunohistochemically quantified fiber type area with the myosin heavy chain (MyHC) contents of a bundle of fibers from a human masticatory muscle. The total cross-sectional areas were determined immunohistochemically for the three major fiber types (I, IIA, and IIX) in bundles of fibers (n=42) taken from the anterior and posterior belly of the human digastric muscle (n=7). The relative MyHC contents of the same fiber bundles were determined electrophoretically (MyHC-I, -IIA, and -IIX; anterior, 32%, 35%, and 33%; posterior, 39%, 42%, and 19%) and compared with the immunohistochemical data (MyHC-I, -IIA, and -IIX; anterior, 32%, 31%, and 37%; posterior, 39%, 45%, and 15%). No significant differences were seen in the mean fiber type distribution between the two techniques; the correlation coefficient ranged from 0.71 to 0.96. The correlation coefficient was higher for MyHC type I and MyHC type IIX than for MyHC type IIA. The MyHC contents of single fibers taken from the posterior belly indicated that many fibers in this belly co-express MyHC-IIA and MyHC-IIX. Despite the presence of these hybrid fibers, the correspondence between both methods was relatively large.

A telemetry system to chronically record muscle activity in middle-sized animals.

Radio-telemetry enables the long-term recordings of biopotentials that may be obtained in freely moving animals without interference by the experimenter. The purpose of this study was to test a fully implantable device for: (1) its transmission range; (2) the characteristics of the transmitted signals; and (3) its actual application in long-term in vivo registration of EMG. Transmission range was tested by changing the device's position relative to the receiver. Computer simulation of the filtering characteristics provided comparison of original and transmitted signals. Implantation of the device in masticatory muscles, followed by analysis of telemetred signals and determination of activity levels allowed for examination of daily muscle use. The implant's transmission range covered the cage size for middle-sized animals with a minimum of signal dropouts. Transmitted signals were marked by (partial) loss of frequencies beyond 50 Hz, decreased amplitude and slightly delayed timing relative to original waveforms. Analysis of the transmitted EMG revealed that the device can be used for prolonged in vivo EMG registration, detection of peak activity levels, and the examination of general muscle use by the time spent at different levels of activity.